Metacommunity size influences aquatic community composition in a natural mesocosm landscape

Ecosystems are often arranged in naturally patchy landscapes with habitat patches linked by dispersal of species in a metacommunity. The size of a metacommunity, or number of patches, is predicted to influence community dynamics and therefore the structure and function of local communities. However, such predictions have yet to be experimentally tested using full food webs in natural metacommunities. We used the natural mesocosm system of aquatic macroinvertebrates in bromeliad phytotelmata to test the effect of the number of patches in a metacommunity on species richness, abundance, and community composition. We created metacommunities of varying size using fine mesh cages to enclose a gradient from a single bromeliad up to the full forest. We found that species richness, abundance, and biomass increased from enclosed metacommunities to the full forest size and that diversity and evenness also increased in larger enclosures. Community composition was affected by metacommunity size across the full gradient, with a more even detritivore community in larger metacommunities, and taxonomic groups such as mosquitoes going locally extinct in smaller metacommunities. We were able to divide the effects of metacommunity size into aquatic and terrestrial habitat components and found that the importance of each varied by species; those with simple life cycles were only affected by local aquatic habitat whereas insects with complex life cycles were also affected by the amount of terrestrial matrix. This differential survival of obligate and non‐obligate dispersers allowed us to partition the beta‐diversity between metacommunities among functional groups. Our study is one of the first tests of metacommunity size in a natural metacommunity landscape and shows that both diversity and community composition are significantly affected by metacommunity size. Synthesis Natural food webs are sensitive to meta‐community size, i.e. the number of patches connected through dispersal. We provide an empirical test using the aquatic foodweb associated within bromeliads as a model system. When we reduced the number of bromeliad patches connect through dispersal, we found a clear change of the foodweb in terms of population sizes, beta diversity, community composition and predator‐prey ratios. The response of individual taxa was predictable based on species traits including dispersal modes, life cycle, and adult resource requirements. Our study demonstrates that community structure is strongly influenced by the interplay of species traits and landscape properties.     

Integrating internal and external dispersal in metacommunity assembly: preliminary theoretical analyses

Internal dispersal, which occurs among local communities within a metacommunity, and external dispersal, which supplies immigrants from outside the metacommunity, can both have a major impact on species diversity. However, few studies have considered the two simultaneously. Here I report preliminary computer-simulation results to suggest that internal and external dispersal can interact to influence species richness. Specifically, the results show that internal dispersal did not affect species richness under frequent external dispersal, whereas it enhanced richness in local communities while decreasing richness in metacommunities under infrequent external dispersal. Conversely, external dispersal influenced species richness in local communities more greatly in the absence of internal dispersal than in its presence, while external dispersal did not affect richness in metacommunities regardless of internal dispersal. Furthermore, internal and external dispersal interactively determined the importance of community assembly history in generating and maintaining variation in local community structure. Overall, these results suggest that the two dispersal types can reciprocally provide the context in which each affects species diversity and therefore that their effects cannot be understood in isolation of the other.Electronic Supplementary Material Supplementary material is available for this article at Tadashi Fukami is the recipient of the ninth Denzaburo Miyadi Award.     

Warming induces synchrony and destabilizes experimental pond zooplankton metacommunities

The spatial insurance hypothesis predicts that intermediate rates of dispersal between patches in a metacommunity allow species to track favourable conditions, preserving diversity and stabilizing biomass at local and regional scales. However, theory is unclear as to whether dispersal will provide spatial insurance when environmental conditions are changing directionally. In particular, increased temperatures as a result of climate change are expected to cause synchronous growth or decline across species and communities, and this has the potential to erode the stabilizing compensatory dynamics facilitated by dispersal. Here we report on an experimental test of how dispersal affects the diversity and stability of metacommunities under warming using replicate two‐patch pond zooplankton metacommunities. Initial differences in local community composition and abiotic conditions were established by seeding each patch in the metacommunities with plankton and sediment from one of two natural ponds that differed in water chemistry and species composition. We exposed metacommunities to a 2°C increase in average ambient temperature, crossed with three rates of dispersal (none, intermediate, high). In ambient conditions, intermediate dispersal rates preserved diversity and stabilized metacommunities by promoting spatially asynchronous fluctuations in biomass, especially between local populations of the dominant genus, Ceriodaphnia. However, warming synchronized their populations so that these effects of dispersal were lost. Furthermore, because the stabilizing effect of dispersal was primarily due to asynchronous fluctuations between populations of a single genus, metacommunity biomass was stabilized, but dispersal did not stabilize local community biomass. Our results show that dispersal can preserve diversity and provide stability to metacommunities, but also show that this benefit can be eroded when warming is directional and synchronous across patches of a metacommunity, as is expected with climate warming.     

Modularity along organism dispersal gradients challenges a prevailing view of abrupt transitions in animal landscape perception

A common property of landscapes and metacommunities is the occurrence of abrupt shifts in connectivity along gradients of individual dispersal abilities. Animals with short‐range dispersal capability perceive fragmented landscapes, but organisms moving across critical thresholds perceive continuous landscapes. This qualitative shift in landscape perception may determine several attributes of local communities and the dynamics of whole metacommunities. Modularity describes the existence in some communities of relatively high numbers of mutual connections favoring the movement of neighboring individuals (even when each individual is able to reach any patch in the landscape). Local patch linkages and metacommunity connectivity along gradients of dispersal ability have been reported frequently. However, the intermediate level of structure captured by modularity has not been considered. We evaluated landscape connectivity and modularity along gradients of individual dispersal abilities. Random landscapes with different degrees of cell aggregation and occupancy were simulated; we also analyzed ten real ecosystems. An expected, a shift in landscape connectivity was always detected; modularity consistently decreased gradually along dispersal gradients in both simulated networks and empirical landscapes. Neutral metacommunities within simulated landscapes demonstrated that modularity and connectivity may reflect landscape traits in the shaping of metacommunity diversity. Average beta‐diversity was strongly associated with modularity, particularly with low migration rates, while connectivity trends tracked changes in beta‐diversity at intermediate to high migrations rates. Consequently, while some species are able to perceive abrupt transitions in the landscape, many others probably experience a gradual continuum in landscape perception, contrary to predictions from previous analyses. Furthermore, the gradual behavior of modularity indicates that it may represent an exceptional early‐warning tool that measures system distance to tipping points. Our study highlights the multiple perceptions that different species may have of a single landscape and shows, for the first time, a theoretical and empirical relationship between landscape modularity, and metacommunity diversity.     

Community patterns in source-sink metacommunities

We present a model of a source-sink competitive metacommunity, defined as a regional set of communities in which local diversity is maintained by dispersal. Although the conditions of local and regional coexistence have been well defined in such systems, no study has attempted to provide clear predictions of classical community-wide patterns. Here we provide predictions for species richness, species relative abundances, and community-level functional properties (productivity and space occupation) at the local and regional scales as functions of the proportion of dispersal between communities. Local (alpha) diversity is maximal at an intermediate level of dispersal, whereas between-community (beta) and regional (gamma) diversity decline as dispersal increases because of increased homogenization of the metacommunity. The relationships between local and regional species richness and the species rank abundance distributions are strongly affected by the level of dispersal. Local productivity and space occupation tend to decline as dispersal increases, resulting in either a hump-shaped or a positive relationship between species richness and productivity, depending on the scale considered (local or regional). These effects of dispersal are buffered by decreasing species dispersal success. Our results provide a niche-based alternative to the recent neutral-metacommunity model and have important implications for conservation biology and landscape management.     

Metacommunity Structure of Stream Insects across Three Hierarchical Spatial scales

A major challenge in community ecology is to understand the underlying factors driving metacommunity (i.e., a set of local communities connected through species dispersal) dynamics. However, little is known about the effects of varying spatial scale on the relative importance of environmental and spatial (i.e., dispersal related) factors in shaping metacommunities and on the relevance of different dispersal pathways. Using a hierarchy of insect metacommunities at three spatial scales (a small, within‐stream scale, intermediate, among‐stream scale, and large, among‐sub‐basin scale), we assessed whether the relative importance of environmental and spatial factors shaping metacommunity structure varies predictably across spatial scales, and tested how the importance of different dispersal routes vary across spatial scales. We also studied if different dispersal ability groups differ in the balance between environmental and spatial control. Variation partitioning showed that environmental factors relative to spatial factors were more important for community composition at the within‐stream scale. In contrast, spatial factors (i.e., eigenvectors from Moran's eigenvector maps) relative to environmental factors were more important at the among‐sub‐basin scale. These results indicate that environmental filtering is likely to be more important at the smallest scale with highest connectivity, while dispersal limitation seems to be more important at the largest scale with lowest connectivity. Community variation at the among‐stream and among‐sub‐basin scales were strongly explained by geographical and topographical distances, indicating that overland pathways might be the main dispersal route at the larger scales among more isolated sites. The relative effect of environmental and spatial factors on insect communities varied between low and high dispersal ability groups; this variation was inconsistent among three hierarchical scales. In sum, our study indicates that spatial scale, connectivity, and dispersal ability jointly shape stream metacommunities.     

Climate change in metacommunities: dispersal gives double-sided effects on persistence

Climate change is increasingly affecting the structure and dynamics of ecological communities both at local and at regional scales, and this can be expected to have important consequences for their robustness and long-term persistence. The aim of the present work is to analyse how the spatial structure of the landscape and dispersal patterns of species (dispersal rate and average dispersal distance) affects metacommunity response to two disturbances: (i) increased mortality during dispersal and (ii) local species extinction. We analyse the disturbances both in isolation and in combination. Using a spatially and dynamically explicit metacommunity model, we find that the effect of dispersal on metacommunity persistence is two-sided: on the one hand, high dispersal significantly reduces the risk of bottom-up extinction cascades following the local removal of a species; on the other hand, when dispersal imposes a risk to the dispersing individuals, high dispersal increases extinction risks, especially when dispersal is global. Large-bodied species with long generation times at the highest trophic level are particularly vulnerable to extinction when dispersal involves a risk. This suggests that decreasing the mortality risk of dispersing individuals by improving the quality of the habitat matrix may greatly increase the robustness of metacommunities.     

Towards understanding the organisation of metacommunities in highly dynamic ecological systems

Community ecology recognises today that local biological communities are not only affected by local biotic interactions and abiotic environmental conditions, but also by regional processes (e.g. dispersal). While much is known about how metacommunities are organised in space in terrestrial, marine and freshwater ecological systems, their temporal variations remain poorly studied. Here, we address the question of the dynamics of metacommunities in highly variable systems, using intermittent rivers (IRs), those rivers which temporarily stop flowing or dry up, as a model system. We first review how habitat heterogeneity in space and time influences metacommunity organisation. Second, we compare the metacommunities in IRs to those in perennial rivers (PRs) and develop the idea that IRs could undergo highly dynamic shifts due to the temporal variability in local and regional community processes. Third, we develop the idea that in IRs, metacommunities of the wet and dry phases of IRs are closely intertwined, thereby increasing even more their respective temporal dynamics. Last, we provide a roadmap to stimulate further conceptual and empirical developments of metacommunity research and identify possible applications for improving the management of IRs and other highly dynamic ecological systems. Synthesis Extensive research has examined the importance of local biotic interactions, environmental filtering, and regional processes on community assembly. Movement of organisms between sites, i.e. dispersal, is a major set of processes within this framework. However, subsequent metacommunity organisation also varies over time in ecosystems because habitat characteristics such as configuration and composition continuously shift. Intermittent rivers are an ideal set of systems to examine these ideas because these freshwater systems temporarily cease flowing thereby limiting dispersal events. We proposed the hypothesis that metacommunities in dynamic ecosystems will undergo frequent shifts in structure and composition in response to the temporal variability in environmental filtering and dispersal. In addition to providing a roadmap for developing a more dynamic perspective for community ecology, these framework provides direct insights for the management of intermittent rivers.     

Dispersal and diversity in experimental metacommunities: linking theory and practice

There has been a recent rise in the number of experiments investigating the effect of dispersal on diversity, with many of the predictions for these tests derived from metacommunity theory. Despite the promise of linking observed relationships between dispersal and diversity to underlying metacommunity processes, empirical studies have faced challenges in providing robust tests of theory. We review experimental studies that have tested how dispersal affects metacommunity diversity to determine why shortcomings emerge, and to provide a framework for empirical tests of theory that capture the processes structuring diversity in natural metacommunities. We first summarize recent experimental work to outline trends in results and to highlight common methods that cause a misalignment between empirical studies and the processes described by theory. We then identify the undesired implications of three widely used experimental methods that homogenize metacommunity structure or species traits, and present alternative methods that have been used to successfully integrate experiments and theory in a biologically relevant way. Finally, we present methodological and theoretical insights from three related ecological fields (coexistence, food web and priority effects theory) that, if integrated into metacommunity experiments, could help isolate the independent and joint effects of local interactions and dispersal on diversity, and reveal the mechanisms underlying observed dispersal–diversity patterns. Together, these methods can provide stronger tests of existing theory and stimulate new theoretical explorations. Synthesis Although metacommunity experiments offer a unique opportunity to test classic and emerging theory on the relationship between dispersal and diversity, several common challenges have hindered robust tests of theory. We outline how emerging theory on the invasion criterion, food webs and priority effects could be help clarify when and how dispersal affects metacommunity diversity, and identify when experimental approaches that homogenize metacommunities fail to test existing theory. By forging better links between theoretical and empirical work, we hope to motivate novel and improved experimental approaches to understanding the joint effects of local and regional processes on diversity.     

Spatial averaging and disturbance lead to high productivity in aquatic metacommunities

Dispersal in heterogeneous ecosystems, such as coastal metacommunities, is a major driver of diversity and productivity. According to theory, both species richness and spatial averaging shape a unimodal relationship of productivity with dispersal. We experimentally tested the hypothesis that disturbances acting on local patches would buffer the loss of productivity at high dispersal by preventing synchronized species oscillations. To simulate these disturbances, our experimental assemblages involved species that self‐organized in isolation under three inflow pulsing frequencies, where hydraulic displacement and nutrient loading affected assemblage diversity and composition. At steady‐state, the emerging isolated assemblages were connected at three levels of dispersal creating three metacommunities of different connectivity. Consistent with theory, as dispersal increased, species richness in the metacommunity declined; productivity however remained high. This occurred because the most productive species in our study (which dominated the isolated patch of intermediate inflow pulsing frequency) dominated all three patches (low, intermediate and high inflow pulsing frequencies) after dispersal commenced in our metacommunities. This experimental result provides empirical support for the mechanism of spatial averaging. Furthermore, disturbances, in the form of localized pulsed inflows, prevented population oscillation synchrony caused by homogenization. Overall, our observations suggest that localized environmental fluctuations and the identity of species seem to be more influential than dispersal in shaping the diversity and composition of phytoplankton assemblages and stabilizing productivity.     

Effect of landscape context on fish metacommunity structuring in stream networks

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Dispersal restricts local biomass but promotes the recovery of metacommunities after temperature stress

Landscape connectivity can increase the capacity of communities to maintain their function when environments change by promoting the immigration of species or populations with adapted traits. However, high immigration may also restrict fine tuning of species compositions to local environmental conditions by homogenizing the community. Here we demonstrate that dispersal generates such a tradeoff between maximizing local biomass and the capacity of model periphyton metacommunities to recover after a simulated heat wave. In non‐disturbed metacommunities, dispersal decreased the total biomass by preventing differentiation in species composition between the local patches making up the metacommunity. On the contrary, in metacommunities exposed to a realistic summer heat wave, dispersal promoted recovery by increasing the biomass of heat tolerant species in all local patches. Thus, the heat wave reorganized the species composition of the metacommunities and after an initial decrease in total biomass by 38.7%, dispersal fueled a full recovery of biomass in the restructured metacommunities. Although dispersal may decrease equilibrium biomass, our results highlight that connectivity is a key requirement for the response diversity that allows ecological communities to adapt to climate change through species sorting.     

Effects of total resources, resource ratios, and species richness on algal productivity and evenness at both metacommunity and local scales

The study of the interrelationship between productivity and biodiversity is a major research field in ecology. Theory predicts that if essential resources are heterogeneously distributed across a metacommunity, single species may dominate productivity in individual metacommunity patches, but a mixture of species will maximize productivity across the whole metacommunity. It also predicts that a balanced supply of resources within local patches should favor species coexistence, whereas resource imbalance would favor the dominance of one species. We performed an experiment with five freshwater algal species to study the effects of total supply of resources, their ratios, and species richness on biovolume production and evenness at the scale of both local patches and metacommunities. Generally, algal biovolume increased, whereas algal resource use efficiency (RUE) and evenness decreased with increasing total supply of resources in mixed communities containing all five species. In contrast to predictions for biovolume production, the species mixtures did not outperform all monocultures at the scale of metacommunities. In other words, we observed no general transgressive overyielding. However, RUE was always higher in mixtures than predicted from monocultures, and analyses indicate that resource partitioning or facilitation in mixtures resulted in higher-than-expected productivity at high resource supply. Contrasting our predictions for the local scale, balanced supply of resources did not generally favor higher local evenness, however lowest evenness was confined to patches with the most imbalanced supply. Thus, our study provides mixed support for recent theoretical advancements to understand biodiversity-productivity relationships.     

Salinisation reduces biodiversity in neighbouring freshwater patches in experimental metacommunities

1. Aquatic ecosystems are biodiversity hot spots across many landscapes; therefore, the degradation of these habitats can lead to decreases in biodiversity across multiple scales. Salinisation is a global issue that threatens freshwater ecosystems by reducing water quality and local biodiversity. The effects of salinity on local processes have been studied extensively; however, the effects of salinisation or similar environmental stressors within a metacommunity (a dispersal network of several distinct communities) have not been explored.
2. We tested how the spatial heterogeneity and the environmental contrast between freshwater and saline habitat patches influenced cladoceran biodiversity and species composition at local and regional scales in a metacommunity mesocosm experiment. We defined spatial heterogeneity as the proportion of freshwater to saltwater patches within the metacommunity, ranging from a freshwater-dominated metacommunity to a saltwater-dominated metacommunity. Environmental contrast was defined as the environmental distance between habitat patches along the salinity gradient in which low-contrast metacommunities consisted of freshwater and low-salinity patches and high-contrast metacommunities consisted of freshwater and high-salinity patches.
3. We hypothesised that the α-richness of freshwater patches and metacommunity γ-richness would decrease as freshwater patches became less abundant along the spatial heterogeneity gradient in both low- and high-contrast metacommunities, because there would be fewer freshwater patches that could serve as source populations for declining populations. We hypothesised that low-contrast metacommunities would support more species across the spatial heterogeneity gradient than high-contrast metacommunities, because, via dispersal, low-salinity patches can support halotolerant freshwater species that can mitigate population declines in neighbouring freshwater patches, whereas` high-salinity patches will mostly support halophilic species, providing fewer potential colonisers to freshwater patches.
4. We found that α-richness of freshwater mesocosms and metacommunity γ-richness declined in saline-dominated metacommunities regardless of the environmental contrast between the freshwater and saline mesocosms. We found that environmental contrast influenced freshwater and saline community composition in low-contrast metacommunities by increasing the abundances of species that could tolerate low-salinity environments through dispersal, whereas freshwater and high-salinity communities showed limited interactions through dispersal.
5. Freshwater mesocosms had a disproportionate effect on the local and regional biodiversity in these experimental metacommunities, indicating that habitat identity may be more important than habitat diversity for maintaining biodiversity in some metacommunities. This study further emphasises the importance in maintaining multiple species-rich habitat patches across landscapes, particularly those experiencing landscape-wide habitat degradation.

     

Condition‐dependent movement and dispersal in experimental metacommunities

Dispersal and the underlying movement behaviour are processes of pivotal importance for understanding and predicting metapopulation and metacommunity dynamics. Generally, dispersal decisions are condition‐dependent and rely on information in the broad sense, like the presence of conspecifics. However, studies on metacommunities that include interspecific interactions generally disregard condition‐dependence. Therefore, it remains unclear whether and how dispersal in metacommunities is condition‐dependent and whether rules derived from single‐species contexts can be scaled up to (meta)communities. Using experimental protist metacommunities, we show how dispersal and movement depend on and are adjusted by the strength of interspecific interactions. We found that the predicting movement and dispersal in metacommunities requires knowledge on behavioural responses to intra‐ and interspecific interaction strengths. Consequently, metacommunity dynamics inferred directly from single‐species metapopulations without taking interspecific interactions into account are likely flawed. Our work identifies the significance of condition‐dependence for understanding metacommunity dynamics, stability and the coexistence and distribution of species.     

Dispersal, edaphic fidelity and speciation in species-rich Western Australian shrublands: evaluating a neutral model of biodiversity

Over evolutionary time, the number of species in a community reflects the balance between the rate of speciation and the rate of extinction. Over shorter time‐scales local species richness is also affected by how often species move into and out of the local community. These processes are at the heart of Hubbell's ‘unified neutral theory of biodiversity’ ( Hubbell 2001 ). Hubbell's spatially implicit, dispersal‐limited neutral model is the most widely used of the many implementations of neutral theory and it provides an estimate of the rate of speciation in a metacommunity (if metacommunity size is known) and the rate at which species migrate into the local community from the wider metacommunity. Recently, this neutral model has been used to compare rates of speciation and migration in the species‐rich fynbos of South Africa and in neotropical forests. Here we use new analytical methods for estimating the neutral model's parameters to infer speciation and dispersal rates for three sites in species‐rich sclerophyll shrublands (equivalent to fynbos) in Western Australia (WA). Our estimates suggest that WA shrublands are intermediate between fynbos and tropical rainforest in terms of speciation and dispersal. Although a weak test, the model predicts species abundance distributions and species accumulation curves similar to those observed at the three sites. The neutral model's predictions also remain plausible when confronted with independent data describing: (1) known edaphic relationships between sites, (2) estimates of metacommunity species richness and (3) rates of speciation among resprouters and nonsprouters. Two of the site pairs, however, show species turnovers significantly different from those predicted by the spatially implicit form of the neutral model that we use. This suggests that non‐neutral processes, in this case probably edaphic specialisation, are important in the WA shrubland metacommunity. The neutral model predicts similar rates of speciation in resprouter and sprouter taxa, a finding supported by recent molecular phylogenies. Finally, when converted into temporally scaled speciation rates and species longevities, the estimates produced by the neutral model seem implausible. The apparent departure from neutrality in the turnover of species between some sites and the implausible temporal dynamics may be due to the particular model chosen and does not reduce the significance of our other results, which confirm that local dispersal limitation, coupled with broader scale edaphic fidelity, combine to structure this biodiverse metacommunity.     

A comparative analysis of metacommunity types in the freshwater realm

Most metacommunity studies have taken a direct mechanistic approach, aiming to model the effects of local and regional processes on local communities within a metacommunity. An alternative approach is to focus on emergent patterns at the metacommunity level through applying the elements of metacommunity structure (EMS; Oikos, 97, 2002, 237) analysis. The EMS approach has very rarely been applied in the context of a comparative analysis of metacommunity types of main microbial, plant, and animal groups. Furthermore, to our knowledge, no study has associated metacommunity types with their potential ecological correlates in the freshwater realm. We assembled data for 45 freshwater metacommunities, incorporating biologically highly disparate organismal groups (i.e., bacteria, algae, macrophytes, invertebrates, and fish). We first examined ecological correlates (e.g., matrix properties, beta diversity, and average characteristics of a metacommunity, including body size, trophic group, ecosystem type, life form, and dispersal mode) of the three elements of metacommunity structure (i.e., coherence, turnover, and boundary clumping). Second, based on those three elements, we determined which metacommunity types prevailed in freshwater systems and which ecological correlates best discriminated among the observed metacommunity types. We found that the three elements of metacommunity structure were not strongly related to the ecological correlates, except that turnover was positively related to beta diversity. We observed six metacommunity types. The most common were Clementsian and quasi‐nested metacommunity types, whereas Random, quasi‐Clementsian, Gleasonian, and quasi‐Gleasonian types were less common. These six metacommunity types were best discriminated by beta diversity and the first axis of metacommunity ecological traits, ranging from metacommunities of producer organisms occurring in streams to those of large predatory organisms occurring in lakes. Our results showed that focusing on the emergent properties of multiple metacommunities provides information additional to that obtained in studies examining variation in local community structure within a metacommunity.     

Temporal dynamics of a local fish community are strongly affected by immigration from the surrounding metacommunity

A 5‐year time series of annual censuses was collected from a large floodplain lake to determine how dynamics of the local fish community were affected by changes in hydrological connectivity with the surrounding metacommunity. The lake was disconnected from the metacommunity for 1 year prior to our study and remained disconnected until 3 months before our third annual census, when a flood reconnected the lake to the metacommunity. We determined how changes in connectivity affected temporal dynamics of (1) local community composition and (2) the population composition, condition, and growth of catfish, to shed light on how immigration of other species might affect local population dynamics. Before reconnection, the community was likely shaped by interactions between the local environment and species traits. The reconnection caused significant immigration and change in community composition and correlated with a significant and abrupt decline in catfish condition, growth, and abundance; effects likely due to the immigration of a competitor with a similar trophic niche: carp. The community was slow to return to its preconnection state, which may be due to dispersal traits of the fishes, and a time‐lag in the recovery of the local catfish population following transient intensification of species interactions. The dynamics observed were concordant with the species sorting and mass‐effects perspectives of metacommunity theory. Floods cause episodic dispersal in floodplain fish metacommunities, and so, flood frequency determines the relative importance of regional and local processes. Local processes may be particularly important to certain species, but these species may need sufficient time between floods for population increase, before the next flood‐induced dispersal episode brings competitors and predators that might cause population decline. Accordingly, species coexistence in these metacommunities may be facilitated by spatiotemporal storage effects, which may in turn be regulated by flood frequency.     

The metacommunity concept: a framework for multi-scale community ecology

The metacommunity concept is an important way to think about linkages between different spatial scales in ecology. Here we review current understanding about this concept. We first investigate issues related to its definition as a set of local communities that are linked by dispersal of multiple potentially interacting species. We then identify four paradigms for metacommunities: the patch‐dynamic view, the species‐sorting view, the mass effects view and the neutral view, that each emphasizes different processes of potential importance in metacommunities. These have somewhat distinct intellectual histories and we discuss elements related to their potential future synthesis. We then use this framework to discuss why the concept is useful in modifying existing ecological thinking and illustrate this with a number of both theoretical and empirical examples. As ecologists strive to understand increasingly complex mechanisms and strive to work across multiple scales of spatio‐temporal organization, concepts like the metacommunity can provide important insights that frequently contrast with those that would be obtained with more conventional approaches based on local communities alone.     

Contrasting metacommunity structure and beta diversity in an aquatic‐floodplain system

Habitat connectivity and dispersal interact to structure metacommunities, but few studies have examined these patterns jointly for organisms across the aquatic–terrestrial ecotone. We assessed metacommunity structure and beta diversity patterns of instream benthic invertebrates, riparian carabid beetles (Order: Coleoptera; Family: Carabidae) and riparian spiders (Order: Araneae) at fifteen sites in a river‐floodplain system. Sampling took place over a three‐year period (2010–2012) in the Rhine‐Main‐Observatory LTER site on the Kinzig River, central Germany. This allowed disentangling the combined influence, and temporal variability, of habitat connectivity (i.e. between aquatic and terrestrial) and dispersal ability (i.e. between spiders and beetles, and aerial and aquatic dispersing invertebrates) on the dominant paradigms structuring these metacommunities. We found mostly consistent differences in the manner that metacommunities were structured between groups, with lower levels of variability explained for beetles compared to the other groups. Beetles were consistently structured more by turnover than nestedness components, with greater beta diversity than expected by chance and a minor spatial compared to environmental signal emerging with variance partitioning. Conversely, spiders and benthic invertebrates had lower beta diversity and greater nestedness than null expectation, and a clearer spatial signal controlling metacommunity structure. Our results suggest varying levels of mass effects and species sorting shape river‐floodplain metacommunities, depending on habitat connectivity and dispersal ability. That is, greater connectivity and lower fragmentation along the river compared to the terrestrial zone promoted mass effects, and differences in overall dispersal ability and mode (i.e. active and passive) for instream and riparian communities shifted paradigms between mass effects and species sorting.